Investigating the effect of ballasting by CaCO3 in Emiliania huxleyi, II: Decomposition of particulate organic matter

The quantitative relationship between organic carbon and mineral contents of particles sinking below 1800 m in the ocean indicates that organisms with mineral shells such as coccolithophores are of special importance for transporting carbon into the deep sea. Several hypotheses about the mechanism behind this relationship between minerals and organic matter have been raised, such as mineral protection of organic matter or enhanced sinking rates through ballast addition. We examined organic matter decomposition of calcifying and non-calcifying Emiliania huxleyi cultures in an experiment that allowed aggregation and settling in rotating tanks. Biogenic components such as particulate carbon, particulate nitrogen, particulate volume, pigments, transparent exopolymer particles (TEP), and particulate amino acids in suspended particles and aggregates were followed over a period of 30 d. The overall pattern of decrease in organic matter, the amount of recalcitrant organic matter left after 30 d, and the compositional changes within particulate organic matter indicated that cells without a shell are more subject to loss than calcified cells. It is suggested that biogenic calcite helps in the preservation of particulate organic matter (POM) by offering structural support for organic molecules. Over the course of the experiment, half the particulate organic carbon in both calcifying and non-calcifying cultures was partitioned into aggregates and remained so until the end of the experiment. The partial protection of particulate organic matter from solubilization by biominerals and by aggregation that was observed in our experiment may help explain the robustness of the relationship between organic and mineral matter fluxes in the deep ocean.     

大洋桥石藻Gephyrocapsa oceanica对海水酸化与营养盐胁迫的复合响应

本文选取了大洋桥石藻(Gephyrocapsa oceanica)作为实验对象,通过CO₂加富实验模拟海水酸化环境,分析了氮充足和氮相对不足条件下海洋颗石藻对海水酸化的生理响应。结果发现在两种营养盐状态下,CO₂加富均能一定程度促进大洋桥石藻种群增长与Chl a浓度的增加。对比两种营养盐状态,氮相对不足条件下的大洋桥石藻细胞密度和叶绿素含量均有最显著的提高,表明低营养盐浓度和低的氮磷比可能更有利于大洋桥石藻的生长繁殖。电镜观测结果显示酸化对大洋桥石藻的钙化作用具有显著的负影响,并且在氮相对不足条件下,大洋桥石藻的细胞个体变小及比表面积升高。研究结果表明在未来寡营养的大洋上层水体,大气CO₂浓度升高会对大洋桥石藻的生理功能产生负面影响,但可能刺激大洋桥石藻的生长。     

Spatial and temporal variability in coccolithophore abundance and production of PIC and POC in the NE subarctic Pacific during El Niño (1998), La Niña (1999) and 2000

Seasonal variations in coccolithophore abundance, chlorophyll, nutrients and production of particulate organic and inorganic carbon (POC and PIC) were determined along a coastal to oceanic east-west transect (Line P) culminating at Ocean Station Papa in the northeastern subarctic Pacific between 1998 and 2000. Offshore stations generally exhibited low seasonality in chlorophyll concentrations, with moderate seasonality in POC production. Near shelf stations showed a similar pattern to offshore stations, but were also characterized by sporadic events of higher POC productivity. During the 1998 El Niño, June was characterized by low chlorophyll and POC productivity along the transect, presumably as a result of depleted surface nitrate. In contrast, during the 1999 La Niña, and in 2000, higher POC productivity and surface nitrate occurred along the transect in June. Chlorophyll and POC productivity were similar in late summer in all 3 years. The coccolithophore population was usually numerically dominated by Emiliania huxleyi, particularly in June. Along the transect, abundance of coccolithophores was much higher in June during the 1998 El Niño (mean of 221 cells ml⁻¹) than in the 1999 La Niña (mean of 40 cells ml⁻¹), with their abundance in late summers of both years being very low. Abundances were even higher along the transect in June and the late summer of 2000 with sporadic ‘blooms’ of >1000 cells ml⁻¹ at some stations (cruise averages 395 and 552 cell ml⁻¹, respectively). Production rates of PIC did not consistently correlate with areas of high coccolithophore abundance. PIC production was high (100-250 mg C m⁻² d⁻¹) along the transect during June 1998, and low (1-40 mg C m⁻² d⁻¹) during both winters, June 1999 and during late summers of 1998 and 1999. The year 2000 was more complicated, with high rates of PIC production accompanying high abundance of coccolithophores in late summer, but lower rates of PIC production accompanying high coccolithophore numbers in June. Our data suggest that the abundance of coccolithophores and the production rates of PIC in the subarctic are higher than previously thought. Occasional PIC:POC production ratios of 1 or greater in 1998 and 2000 suggest that coccolithophores in this region could have a significant impact on the efficiency of the biological carbon pump.     

Spatial Variability of Living Coccolithophore Distribution in the Western Subarctic Pacific and Western Bering Sea

Vertical distributions of coccolithophores were observed in the depth range 0–50 m in the western subarctic Pacific and western Bering Sea in summer, 1997. Thirty-five species of coccolithophores were collected. Overall, Emiliania huxleyi var. huxleyi was the most abundant taxon, accounting for 82.8% of all coccolithophores, although it was less abundant in the western Bering Sea. Maximum abundance of this species was found in an area south of 41°N and east of 175°E (Transition Zone) reaching >10,000 cells L⁻¹ in the water column. In addition to this species, Coccolithus pelagicus f. pelagicus, which accounted for 4.2% of the assemblage, was representative of the coccolithophore standing crop in the western part of the subarctic Pacific. Coccolithus pelagicus f. hyalinus was relatively abundant in the Bering Sea, accounting for 2.6% of the assemblage. Coccolithophore standing crops in the top 50 m were high south of 41°N (>241 × 10⁶ cells m⁻²) and east of 170°E (542 × 10⁶ cells m⁻²) where temperatures were higher than 12°C and salinities were greater than 34.2. The lowest standing crop was observed in the Bering Sea and Oyashio areas where temperatures were lower than 6–10°C and salinities were less than 33.0. From the coccolithophore volumes, the calcite stocks in the Transition, Subarctic, and the Bering Sea regions were estimated to be 73.0, 9.7, and 6.9 mg m⁻², respectively, corresponding to calcite fluxes of 3.6, 0.5, and 0.3 mg m⁻²d⁻¹ using Stoke's Law. This revised version was published online in July 2006 with corrections to the Cover Date.     

Investigating the effect of ballasting by CaCO3 in Emiliania huxleyi: I. Formation, settling velocities and physical properties of aggregates

To investigate the role of ballasting by biogenic minerals in the export of organic matter in the ocean, a laboratory experiment was conducted comparing aggregate formation and settling velocity of non-calcifying and calcifying strains of the coccolithophore Emiliania huxleyi. Experiments were conducted by making aggregates using a roller table and following aggregate properties during incubation for a period of 40 days. Size, shape, and settling velocities of aggregates were described by image analysis of video pictures recorded during the roller tank incubation. Our results show that biogenic calcite has a strong effect on the formation rate and abundance of aggregates and on aggregate properties such as size, excess density, porosity, and settling velocity. Aggregates of calcifying cells (AGG_CAL) formed faster, were smaller and had higher settling velocities, excess densities, and mass than those of non-calcifying cells (AGG_NCAL). AGG_CAL showed no loss during the duration of the experiment, whereas AGG_NCAL decreased in size after 1 month of incubation. Potential mechanisms that can explain the different patterns in aggregate formation are discussed. Comparison of settling velocities of AGG_CAL and AGG_NCAL with aggregates formed by diatoms furthermore indicated that the ballast effect of calcite is greater than that of opal. Together these results help to better understand why calcite is of major importance for organic matter fluxes to the deep ocean.     

Biogeochemistry and carbon mass balance of a coccolithophore bloom in the northern Bay of Biscay (June 2006)

Primary production (PP), calcification (CAL), bacterial production (BP) and dark community respiration (DCR) were measured along with a set of various biogeochemical variables, in early June 2006, at several stations at the shelf break of the northern Bay of Biscay. The cruise was carried out after the main spring diatom bloom that, based on the analysis of a time-series of remotely sensed chlorophyll-a (Chl-a), peaked in mid-April. Remotely sensed sea surface temperature (SST) indicated the occurrence of enhanced vertical mixing (due to internal tides) at the continental slope, while adjacent waters on the continental shelf were stratified, as confirmed by vertical profiles of temperature acquired during the cruise. The surface layer of the stratified water masses (on the continental shelf) was depleted of inorganic nutrients. Dissolved silicate (DSi) levels probably did not allow significant diatom development. We hypothesize that mixing at the continental slope allowed the injection of inorganic nutrients that triggered the blooming of mixed phytoplanktonic communities dominated by coccolithophores (Emiliania huxleyi) that were favoured with regards to diatoms due to the low DSi levels. Based on this conceptual frame, we used an indicator of vertical stratification to classify the different sampled stations, and to reconstruct the possible evolution of the bloom from the onset at the continental slope (triggered by vertical mixing) through its development as the water mass was advected on-shelf and stratified. We also established a carbon mass balance at each station by integrating in the photic layer PP, CAL and DCR. This allowed computation at each station of the contribution of PP, CAL and DCR to CO₂ fluxes in the photic layer, and how they changed from one station to another along the sequence of bloom development (as traced by the stratification indicator). This also showed a shift from net autotrophy to net heterotrophy as the water mass aged (stratified), and suggested the importance of extracellular production of carbon to sustain the bacterial demand in the photic and aphotic layers.     

Coccolithophores from the central Arabian Sea: Sediment trap results

Sediment trap samples collected from a depth of 1018 m in the Central Arabian Sea Trap (CAST) at 14°28.2′N, 64°35.8′E were analyzed for temporal variation of coccolithophore fluxes from October 1993 to August 1994. Out of the twenty species of coccolithophores encountered,Gephyrocapsa oceanica, Emiliania huxleyi, Umbilicosphaera sibogae andUmbellosphaera irregularis were the most abundant. The total coccolithophore fluxes ranged from 28.5 × 10⁶m^-2d^-1 to 50.3 × 10⁶m^-2d^-1 showing seasonality with higher fluxes during the northeast (NE) monsoon and lower fluxes during the spring intermonsoon. The higher fluxes were attributed to the enhancement of primary production in the central Arabian Sea due to southward extent of nutrients from the northeast Arabian Sea by the prevailing surface currents. Similarly, the occurrences of relatively lower coccolithophore fluxes during the spring intermonsoon and southwest (SW) monsoon were attributed to the low nutrients in the warm, shallow surface mixed layer and downwelling to the south of Findlater Jet respectively in the central Arabian Sea. Some of the coccolithophore species such asE. huxleyi, G. oceanica, Calcidiscus leptoporus andUmbellosphaera tenuis showed signs of dissolution.